Behavioural responses to anthropogenic disturbances

When a habitat undergoes change, the first response of an individual is often behavioural adjustment. This immediate response can determine whether the population will survive or not, as behavioural flexibility gives time for genetic changes to arise later on. Habitat changes that alter reproductive behaviours can have long-lasting effects on populations. If the selective regime has changed under the new conditions, mate choice cues may no longer reliably reflect an individual s quality. Thus, animals have to be able to adjust their reproductive behaviours to the local conditions. The aim of my thesis was to discuss if and how animals are able to respond to rapid anthropogenic environmental change, and to study the mechanisms of the responses and the evolutionary consequences. The main focus was on the effects of human-induced eutrophication on the reproductive behaviour of fishes. Eutrophication is the result of increased nutrient input and can cause dense underwater vegetation and algal blooms. I used fishes from two very different ecosystems as model species, the Baltic Sea threespine stickleback (Gasterosteus aculeatus) and the desert goby (Chlamydogobius eremius), an endemic species of the Lake Eyre region in Central Australia. I investigated the effects of increased habitat complexity on courtship behaviour and the possibility of local differentiation in courtship and nest building behaviour depending on the level eutrophication in the habitat of origin. Furthermore, I observed the effect of turbidity on stickleback nest building behaviour. The results show that threespine stickleback males, which were born in areas that have been eutrophied for decades, court females at a higher intensity than males from clear water areas. Similarly, male desert gobies increased their courtship effort in dense vegetation. Intense courtship could be an adjustment to reduced visibility and lowered predation risk in the densely vegetated sites. However, there were no clear differences in nest building between males from clear and eutrophied areas under standardized conditions. This was expected as Baltic Sea sticklebacks prefer to nest under vegetation cover and are fairly rigid in adjusting their nest characteristics. Nest building was affected by increased turbidity: males built smaller nests with a larger nest entrance in turbid water. The large variation in the magnitude of phytoplankton blooms may require a rapid adjustment of the optimal nest structure to the current conditions. This thesis highlights the complex interactions that are set- off by human-induced changes in habitats and are followed by the immediate behavioural responses. It also encourages more research to tease apart the phenotypic and genetic components of the observed behavioural differences.

Behavioural and morphological dimorphism of the sexes: an account of two primitively eusocial wasps

In social Hymenoptera, foraging, nest building, brood care and all other colony maintenance functions are carried out by the females while males function solely as reproductives. This asymmetry in social roles of the two sexes has led social insect researchers to focus almost exclusively on the females whereas males have remained relatively neglected. We studied two sympatric, primitively eusocial wasps, Ropalidia marginata and Ropalidia cyathiformis, and compared the morphological and behavioural profiles of males and females. Males of both species are smaller in size and weigh less compared to females. Males of the two species live in the nest for different durations. Borrowing from the ecological literature we use novel methods to compute and compare behavioural diversity and behavioural richness and show that females of both species are behaviourally richer and more diverse than the males.

Cannibalism can drive the evolution of behavioural phase polyphenism in locusts

During outbreaks, locust swarms can contain millions of insects travelling thousands of kilometers while devastating vegetation and crops. Such large-scale spatial organization is preceded locally by a dramatic density-dependent phenotypic transition in multiple traits. Behaviourally, low-density solitarious individuals avoid contact with one another; above a critical local density, they undergo a rapid behavioural transition to the gregarious phase whereby they exhibit mutual attraction. Although proximate causes of this phase polyphenism have been widely studied, the ultimate driving factors remain unclear. Using an individual-based evolutionary model, we reveal that cannibalism, a striking feature of locust ecology, could lead to the evolution of density-dependent behavioural phase-change in juvenile locusts. We show that this behavioural strategy minimizes risk associated with cannibalistic interactions and may account for the empirically observed persistence of locust groups during outbreaks. Our results provide a parsimonious explanation for the evolution of behavioural plasticity in locusts.

The energy costs of sexual dimorphism in mole-rats are morphological not behavioural.

Different reproductive strategies of males and females may lead to the evolution of differences in their energetic costs of reproduction, overall energetic requirements and physiological performances. Sexual dimorphism is often associated with costly behaviours (e.g. large males might have a competitive advantage in fighting, which is energetically expensive). However, few studies of mammals have directly compared the energy costs of reproductive activities between sexes. We compared the daily energy expenditure (DEE) and resting metabolic rate (RMR) of males and females of two species of mole-rat, Bathyergus janetta and Georychus capensis (the former is sexually dimorphic in body size and the latter is not) during a period of intense digging when males seek females. We hypothesized that large body size might be indicative of greater digging or fighting capabilities, and hence greater mass-independent DEE values in males of the sexually dimorphic species. In contrast to this prediction, although absolute values of DEE were greater in B. janetta males, mass-independent values were not. No differences were apparent between sexes in G. capensis. By comparison, although RMR values were greater in B. janetta than G. capensis, no differences were apparent between the sexes for either species. The energy cost of dimorphism is most likely to be the cost of maintenance of a large body size, and not the cost of behaviours performed when an individual is large.

Recent advances in the analysis of behavioural organization and interpretation as indicators of animal welfare

While the incorporation of mathematical and engineering methods has greatly advanced in other areas of the life sciences, they have been under-utilized in the field of animal welfare. Exceptions are beginning to emerge and share a common motivation to quantify 'hidden' aspects in the structure of the behaviour of an individual, or group of animals. Such analyses have the potential to quantify behavioural markers of pain and stress and quantify abnormal behaviour objectively. This review seeks to explore the scope of such analytical methods as behavioural indicators of welfare. We outline four classes of analyses that can be used to quantify aspects of behavioural organization. The underlying principles, possible applications and limitations are described for: fractal analysis, temporal methods, social network analysis, and agent-based modelling and simulation. We hope to encourage further application of analyses of behavioural organization by highlighting potential applications in the assessment of animal welfare, and increasing awareness of the scope for the development of new mathematical methods in this area.

Behavioural consequences of imagining intergroup contact with stigmatized outgroups

We investigated whether imagining contact with an outgroup member would change intergroup behaviour. Participants who had imagined a positive interaction with an outgroup member or an unspecified stranger were told that they were about to take part in a discussion task with an outgroup member. They were taken to a room and asked to set out two chairs ready for the discussion while the experimenter left, ostensibly to find the other participant. The distance between the two chairs was then measured. Undergraduate students who imagined talking to an obese individual (Experiment 1) or a Muslim individual (Experiment 2) placed the chairs significantly closer than those in the control condition. They also reported more positive feelings and beliefs regarding Muslims. These findings highlight an important practical application of imagined contact: preparing people for successful face-to-face contact.

Is the Schwabe organ a retained larval eye? Anatomical and behavioural studies of a novel sense organ in adult leptochiton asellus (Mollusca, Polyplacophora) indicate links to larval photoreceptors

The discovery of a sensory organ, the Schwabe organ, was recently reported as a unifying feature of chitons in the order Lepidopleurida. It is a patch of pigmented tissue located on the roof of the pallial cavity, beneath the velum on either side of the mouth. The epithelium is densely innervated and contains two types of potential sensory cells. As the function of the Schwabe organ remains unknown, we have taken a cross-disciplinary approach, using anatomical, histological and behavioural techniques to understand it. In general, the pigmentation that characterises this sensory structure gradually fades after death; however, one particular concentrated pigment dot persists. This dot is positionally homologous to the larval eye in chiton trochophores, found in the same neuroanatomical location, and furthermore the metamorphic migration of the larval eye is ventral in species known to possess Schwabe organs. Here we report the presence of a discrete subsurface epithelial structure in the region of the Schwabe organ in Leptochiton asellus that histologically resembles the chiton larval eye. Behavioural experiments demonstrate that Leptochiton asellus with intact Schwabe organs actively avoid an upwelling light source, while Leptochiton asellus with surgically ablated Schwabe organs and a control species lacking the organ (members of the other extant order, Chitonida) do not (Kruskal-Wallis, H = 24.82, df = 3, p < 0.0001). We propose that the Schwabe organ represents the adult expression of the chiton larval eye, being retained and elaborated in adult lepidopleurans.

Does subjective rating reflect behavioural coding? Personality in 2 month-old dog puppies: An open-field test and adjective-based questionnaire

A number of studies have recently investigated personality traits in non-human species, with the dog gaining popularity as a subject species for research in this area. Recent research has shown the consistency of personality traits across both context and time for adult dogs, both when using questionnaire based methods of investigation and behavioural analyses of the dogs' behaviour. However, only a few studies have assessed the correspondence between these two methods, with results varying considerably across studies. Furthermore, most studies have focused on adult dogs, despite the fact that an understanding of personality traits in young puppies may be important for research focusing on the genetic basis of personality traits. In the current study, we sought to evaluate the correspondence between a questionnaire based method and the in depth analyses of the behaviour of 2-month old puppies in an open-field test in which a number of both social and non-social stimuli were presented to the subjects. We further evaluated consistency of traits over time by re-testing a subset of puppies. The correspondence between methods was high and test-retest consistency (for the main trait) was also good using both evaluation methods. Results showed clear factors referring to the two main personality traits 'extroversion,' (i.e. the enthusiastic, exuberant approach to the stimuli) and 'neuroticism,' (i.e. the more cautious and fearful approach to the stimuli), potentially similar to the shyness-boldness dimension found in previous studies. Furthermore, both methods identified an 'amicability' dimension, expressing the positive interactions the pups directed at the humans stranger, and a 'reservedness' dimension which identified pups who largely chose not to interact with the stimuli, and were defined as quiet and not nosey in the questionnaire.

Cues to action : do they result in belief and behavioural change in women?

With incidence rates of osteoporosis increasing (Osteoporosis Canada, 2007), preventative efforts to minimize costs associated with condition diagnosis are a public health priority. Cues to action are specific internal (e.g., physical symptoms, family member with a condition) or external stimuli (e.g., public service announcements, health education campaigns) that are necessary to trigger appropriate health behaviours and serve to create an awareness of the health threat (Mattson, 1999). To date, limited understanding of the scope of influence cues to action have on health beliefs and behaviour associated with osteoporosis is known. The present investigation was designed to address this gap in the literature. More specifically, the influence of cues to action, a public service announcement (PSA) developed by Osteoporosis Canada and a bone screening by way of Quantitative Ultrasound, on health beliefs and health-enhancing physical activity (HEPA) across a four week period was investigated. Peri-and postmenopausal women (N= 174) were randomly assigned to one of three conditions 1) an osteoporosis public service announcement (PSA) condition; 2) a bone screening condition via quantitative ultrasound techniques, and 3) a PSA attention control condition. Health beliefs associated with osteoporosis were taken at three time points: prior to the cue to action intervention, immediately following the intervention, and four weeks post intervention. Knowledge of osteorporosis risk factors and HEP A were assessed pre and post-intervention only. Results of a regression analysis suggested that baseline health beliefs predicted baseline HEPA (R2 adj = .24; F (9, 161) = 6.49,p = .000; 95% CI = .12 - .35) with exercise barriers (p = -.33) being a negative predictor and health motivation (p = .21) being a positive predictor of HEP A. Baseline health beliefs predicted With incidence rates of osteoporosis increasing (Osteoporosis Canada, 2007), preventative efforts to minimize costs associated with condition diagnosis are a public health priority. Cues to action are specific internal (e.g., physical symptoms, family member with a condition) or external stimuli (e.g., public service announcements, health education campaigns) that are necessary to trigger appropriate health behaviours and serve to create an awareness of the health threat (Mattson, 1999). To date, limited understanding of the scope of influence cues to action have on health beliefs and behaviour associated with osteoporosis is known. The present investigation was designed to address this gap in the literature. More specifically, the influence of cues to action, a public service announcement (PSA) developed by Osteoporosis Canada and a bone screening by way of Quantitative Ultrasound, on health beliefs and health-enhancing physical activity (HEPA) across a four week period was investigated. Peri-and postmenopausal women (N= 174) were randomly assigned to one of three conditions 1) an osteoporosis public service announcement (PSA) condition; 2) a bone screening condition via quantitative ultrasound techniques, and 3) a PSA attention control condition. Health beliefs associated with osteoporosis were taken at three time points: prior to the cue to action intervention, immediately following the intervention, and four weeks post intervention. Knowledge of osteorporosis risk factors and HEP A were assessed pre and post-intervention only. Results of a regression analysis suggested that baseline health beliefs predicted baseline HEPA (R2 adj = .24; F (9, 161) = 6.49,p = .000; 95% CI = .12 - .35) with exercise barriers (p = -.33) being a negative predictor and health motivation (p = .21) being a positive predictor of HEP A. Baseline health beliefs predicted

Level and precision of behavioural thermoregulation in the bearded dragon, Pogona vitticeps : effects of hypoxia and environmental thermal quality /

Most metabolic functions are optimized within a narrow range of body temperatures, which is why thermoregulation is of great importance for the survival and overall fitness of an animal. It has been proposed that lizards will thermoregulate less precisely in low thermal quality environments, where the costs associated with thermoregulation are high; in the case of lizards, whose thermoregulation is mainly behavioural, the primary costs ofthermoregulation are those derived from locomotion. Decreasing thermoregulatory precision in costly situations is a strategy that enhances fitness by allowing lizards to be more flexible to changing environmental conditions. It allows animals to maximize the benefits of maintaining a relatively high body temperature while minimizing energy expenditure. In situations where oxygen concentration is low, the costs of thermoregulation are relatively high (i.e. in relation to the amount of oxygen available for metabolic functions). As a result, it is likely that exposures to hypoxic conditions induce a decrease in the precision of thermoregulation. This study evaluated the effects of hypoxia and low environmental thermal quality, two energetically costly conditions, on the precision and level of thermoregulation in the bearded dragon, Pogona vitticeps, in an electronic temperature-choice shuttle box. Four levels of hypoxia (1O, 7, 5 and 4% 02) were tested. Environmental thermal quality was manipulated by varying the rate of temperature change (oTa) in an electronic temperature-choice shuttle box. Higher oT a's translate into more thermally challenging environments, since under these conditions the animals are forced to move a greater number of times (and hence invest more energy in locomotion) to maintain similar temperatures than at lower oTa's. In addition, lizards were tested in an "extreme temperatures" treatment during which air temperatures of the hot and cold compartments of the shuttle box were maintained at a constant 50 and 15°C respectively. This was considered the most thermally challenging environment. The selected ambient (T a) and internal body temperatures (Tb) of bearded dragons, as well as the thermoregulatory precision (measured by the central 68% ofthe Ta and T b distribution) were evaluated. The thermoregulatory response was similar to both conditions. A significant increase in the size of the Tb range, reflecting a decrease in thermoregulatory precision, and a drop in preferred body temperature of ~2 °C, were observed at both 4% oxygen and at the environment of lowest thermal quality. The present study suggests that in energetically costly situations, such as the ones tested in this study, the bearded dragon reduces energy expenditure by decreasing preferred body temperature and minimizing locomotion, at the expense of precise behavioural thermoregulation. The close similarity of the behavioural thermoregulatory response to two very different stimuli suggests a possible common mechanism and neuronal pathway to the thermoregulatory response.

Behavioural and neural correlates of operant conditioning in Lymnaea stagnalis : role of previous experience during development /

The freshwater mollusc Lymnaea stagnalis was utilized in this study to further the understanding of how network properties change as a result of associative learning, and to determine whether or not this plasticity is dependent on previous experience during development. The respiratory and neural correlates of operant conditioning were first determined in normally reared Lymnaea. The same procedure was then applied to differentially reared Lymnaea, that is, animals that had never experienced aerial respiration during their development. The aim was to determine whether these animals would demonstrate the same responses to the training paradigm. In normally reared animals, a behavioural reduction in aerial respiration was accompanied by numerous changes within the neural network. Specifically, I provide evidence of changes at the level of the respiratory central pattern generator and the motor output. In the differentially reared animals, there was little behavioural data to suggest learning and memory. There were, however, significant differences in the network parameters, similar to those observed in normally reared animals. This demonstrated an effect of operant conditioning on differentially reared animals. In this thesis, I have identified additional correlates of operant conditioning in normally reared animals and provide evidence of associative learning in differentially reared animals. I conclude plasticity is not dependent on previous experience, but is rather ontogenetically programmed within the neural network.

Seasonal changes in behavioural and thermoregulatory responses to hypoxia in the Eastern Chipmunk (Tamias striatus) /

Mammalian heterotherms, such as hibemators, are known to be more tolerant of low oxygen tensions than their homeothermic counterparts. It has been suggested that this relative hypoxia tolerance is related to their ability to deal with dramatic changes in body temperature during entry to and arousal from torpor. However, hibemators demonstrate dramatic seasonality in both daily heterothermy and overall torpor expression. It was of interest to test if seasonal comparisons of normothermic individuals within a single species with the capacity to hibernate produce changes in the response to hypoxia that would reflect a seasonal change in tolerance to low oxygen. In particular, the species studied, the Eastern chipmunk {Tamias striatus), is known to enter into torpor exclusively in the winter. To test for seasonal differences in the metabolic and thermoregulatory responses to hypoxia, flow-through respirometry was used to compare metabolic rate, minimum thermal conductance, body temperature, and a thermal gradient used to assess selected ambient temperature in response to hypoxia in both summer and winter acclimated animals. Although the animals periodically expressed torpor throughout the winter, no differences between season in resting metabolic rate, body temperature or minimum thermal conductance were observed in normoxia. The metabolic trials indicated that chipmunks are less responsive to hypoxia in the winter than they are in the summer. Although body temperature dropped in response to hypoxia in both seasons, the decrease was less in the winter, and there was no corresponding decrease in metabolic rate. Providing the animals with a choice of ambient temperatures in hypoxia resulted in a blunting of the drop in body temperature in both seasons, suggesting that the reported fall in body temperature set point in hypoxia is not fully manifested in the behavioural pathways responsible for thermoregulation in chipmunks. Instead, body temperature in hypoxia appears to be highly dependent on ambient temperature and oxygen concentration. The results of this study suggest that the season in which the responses to hypoxia are measured is important, especially in a heterotherm where seasonality can affect the degree to 1 which the animal is tolerant of hypoxia. Winter-acclimated chipmunks appear more capable of defending metabolic heat production in hypoxia, a response consistent with the increased thermogenic capacity observed in animals that must periodically enter and arouse from torpor during hibernation.

Behavioural characteristics in phylogenetics : a case study using black fly (Diptera: Simuliidae) cocoon spinning behaviour

1-1 is torically, the predominan t method of reconstructing phylogenies has been through the use of morphological characters. There are new techniques now gaining acceptance, including molecular techniques al1d chromosomal information. Altl10ugh the study of behaviour has been used in a comparative framework, these analyses have, historically, been based on intuition. Hennig (1966) devised a neV\' method of reconstructing phylogenies which provided a 110ncircular method for formulating, testing and refining phylogenies. Subsequent s)Tstematists had virtually abandoned ecological and beha\lioural data as primary indicators of phylogenetic relationships (Brooks and McLennan 1991). Therefore, in a modern cladistic framework (sensu Hennig) the analysis of behavioural traits remains underrepresented as a method of reconstructing phylogenies. This thesis will reconstruct the phylogeny for species of black flies (Diptera: Simuliidae), using two steps. The first step is to thoroughl)' understand and explain the cocoon spinning in black fly larvae. There have bee115 previous descriptions of cocoon spinning, but all were incomplete or erroneous. The advances in technology, including video recorders and VCRs, have allowed this behaviour to be analyzed in great detail in 20 different species. A complete description of the cocoon spinning of Simulium \littatum is given. This description will be used as a template for the other species observed. The description and understanding of cococ)n spinning was the first step in undertaking a phylogenetic analysis using this behaviour. The behaviour was then broken down and analyzed, revealing 23 characters, 3 either qualitative and quantitative in nature. These characters were assessed in a cladistic framework (sensu Hennig) and a phylogenetic tree was reconstructed with a e.I of 0.91 and an R.I. of 0.96. This phylogenetic tree closely resembles a previously established pllylogenetic tree produced from morphological and cytological information. The importance of this result is the indication that, contrary to some authors, behavioural characters, if used properly, can add very informative characters to a data set.

Behavioural, mensural, and ecological correlates of polygyny in the eastern meadowlark (Sturnella magna)

During 1982 and 1983 I studied male attributes and attributes of the territory of male Eastern Meadowlarks (Sturnella magna) in order to determine whether there was a correlation between any of the attributes investigated and the number of females attracted by a male. Seventeen males, nine of which were polygynous and eight monogamous, were studied in 1982 and sixteen males.of which .. seven were polygynous and nine ~onogamous, were studied in 1983. The study was conducted in Short Hills Park, 10 km southwest of St. Catharines, Ontario and was designed to compare two hypotheses: the "sexy son" hypothesis (Weatherhead and Robertson,1977) and the polygyny threshold model (Verner and Willson,1966, Orians, 1969). Male attributes investigated were male size and song behaviour. Six measures of male size were taken: weight, flattened and natural wing chord length, culmen length, bill depth and length of the tarsometatarsus. In 1983 song repertoire size and song versatility measures were investigated. Attributes of the territory studied were: territory size, density of plant stems, percentage plant cover and measures of vegetation structure. In 1983 Arthropods were collected from each territory and sorted according to taxonomic group and size. During 1983, territory attributes were sampled twice, once early and once later in the nesting season. Analysis of data involved univariate comparisons between monogamous and polygynous males using T-tests and multivariate comparisons were made using discriminant function analysis (DFA) and principle components analysis (PCA).No correlations were found between the number of females attracted with, .ny measure of male size or with me, .sures of song versatili or size of song repertoire. Also no correlation was found between terri size and the number of females nesting on a terri . Some attributes of the male's terri id distinguish between monog,mous and po s males of thistudy. Analysis of Arthropod numbers showed that e~ .eran counts were significantly great~r on polygynous territories, a1 the total numb~rs of Arthropods collected showed no s fico .nt differences between territories of monogamous and po males. DFA chose ear teran and Hymenopteran counts as multivariate discriminators; both variables we' e more vegetation revealed that there were no univariate differences between the two groups of males fOT 1982 stem densities, but ~ spp. and Solidago spp. were chosen DFA as multivariate discriminators. The total number of plant stems and of Vicia spp. stems were s ficantly the early 1983 ing on monogamous territories for however DFA found no multivariate discriminators" Variables concerned with the overall aspects of vegetation structure showed significant differences between territories of monogamous and polygynous males. DFA of the 1982 sampling of vegetation structure showed significantly greater mat depth and vegetation height on polygynous territories, a finding which was not supported, however, by peA. For the early 1983 sampling period, plant height was greater on polygynous territories. Multivariate analysis identified greater green cover on polygynous territories, greater ground cover on monogamous territories, and greater depth of mat material on monogamous territories as discriminators between territories of monogamous and polygynous males. A DFA on the major variables of the study showed no significant difference between the territories of monogamous and polygynous male Meadowlarks. Of the correlations found, some were for non-prey Arthr~ods, for cover plants with very small samples sizes, or for variables which were greater for monogamous males during one sampling period and polygynous males during the next. While multivariate discriminators were found, peA showed no grouping of monogamous or polygynous males according to any of the variables investigated. On the basis of the univariate and multivariate analysis of major variables, I concluded that there were no correlations between the number of females attracted with male attributes and no unambiguous correlation with attributes of the territory. My study does not unequivocally support either the "sexy son" or the polygyny threshold hypothesis.